Homology of arthropod anterior appendages revealed by Hox gene expression in a sea spider

Arthropod head segments offer a paradigm for understanding the diversification of form during evolution, as a variety of morphologically diverse appendages have arisen from them. There has been long-running controversy, however, concerning which head appendages are homologous among arthropods, and from which ancestral arrangement they have been derived. This controversy has recently been rekindled by the proposition that the probable ancestral arrangement, with appendages on the first head segment, has not been lost in all extant arthropods as previously thought, but has been retained in the pycnogonids, or sea spiders. This proposal was based on the neuroanatomical analysis of larvae from the sea spider Anoplodactylus sp., and suggested that the most anterior pair of appendages, the chelifores, are innervated from the first part of the brain, the protocerebrum. Our examination of Hox gene expression in another sea spider, Endeis spinosa, refutes this hypothesis. The anterior boundaries of Hox gene expression domains place the chelifore appendages as clearly belonging to the second head segment, innervated from the second part of the brain, the deutocerebrum. The deutocerebrum must have been secondarily displaced towards the protocerebrum in pycnogonid ancestors. As anterior-most appendages are also deutocerebral in the other two arthropod groups, the Euchelicerata and the Mandibulata, we conclude that the protocerebral appendages have been lost in all extant arthropods.     

A palaeontological solution to the arthropod head problem

The composition of the arthropod head has been one of the most controversial topics in zoology, with a large number of theories being proposed to account for it over the last century. Although fossils have been recognized as being of potential importance in resolving the issue, a lack of consensus over their systematics has obscured their contribution. Here, I show that a group of previously problematic Cambrian arthropods from the Burgess Shale and Chengjiang faunas form a clade close to crown-group euarthropods, the group containing myriapods, chelicerates, insects and crustaceans. They are characterized by modified or even absent endopods, and two pre-oral appendages. Comparison with reconstructions of the crown-group euarthropod ground plan and recent investigations into onychophorans demonstrates that these two appendages are the first antenna (of extant crustaceans) and a more anterior appendage associated with an ocular segment. The latter appendage has been reduced in all crown-group euarthropods. Its most likely relic is as a component of the labrum. These fossils thus tie together results from disparate living groups (onychophorans and euarthropods).     

A Silurian sea spider

Pycnogonids (sea spiders) are marine arthropods numbering some 1,160 extant species. They are globally distributed in depths of up to 6,000 metres, and locally abundant; however, their typically delicate form and non-biomineralized cuticle has resulted in an extremely sparse fossil record that is not accepted universally. There are two opposing views of their phylogenetic position: either within Chelicerata as sister group to the euchelicerates, or as a sister taxon to all other euarthropods. The Silurian Herefordshire Konservat-Lagerstatte in England (approximately 425 million years (Myr) bp) yields exceptionally preserved three-dimensional fossils that provide unrivalled insights into the palaeobiology of a variety of invertebrates. The fossils are preserved as calcitic void in-fills in carbonate concretions within a volcaniclastic horizon, and are reconstructed digitally. Here we describe a new pycnogonid from this deposit, which is the oldest adult sea spider by approximately 35 Myr and the most completely known fossil species. The large chelate first appendage is consistent with a chelicerate affinity for the pycnogonids. Cladistic analyses place the new species near the base of the pycnogonid crown group, implying that the latter had arisen by the Silurian period.     

Phylogeny and life habits of Early Arthropods-Predation in the Early Cambrian Sea

We investigated two new arthropods from the Maotianshan-Shale fauna of southern China in the course of our research on life strategies, particularly predation, in Early Cambrian marine macrofaunal biota. One form clearly belongs to the so-called "great-appendage" arthropods, animals that were, most likely, active predators catching prey with their first pair of large, specialized frontoventral appendages. Based on this, we hypothesize that the new species and many others, if not all, of the "great-appendage" arthropods were derivatives of the chelicerate stem lineage and not forms having branched off at different nodes along the evolutionary lineage of the Arthropoda. Rather, we consider the "great-appendage" arthropods as belonging to a monophyletic clade, which modified autapomorphically their first pair of appendages (antennae in general arthropod terminology) into raptorial organs for food capture. The second new form resembles another Maotianshan-Shale arthropod, Fuxianhuia protensa, in sharing a head made of only two separate segments, a small segment bearing oval eyes laterally, and another bearing a large tergite, which forms a wide shield freely overhanging the subsequent narrow trunk segments. This segment bears a single pair of rather short anteriorly directed uniramous appendages, considered as the "still" limb-shaped antennae. Particularly the evolutionary status of head and limbs of these two forms suggests that both are representatives of the early part of the stem lineage toward the crown-group of Arthropoda, the Euarthropoda. These forms appear rather unspecialized, but may have been but simple predators. This adds to our hypothesis that predation was a common, if not dominant feeding strategy in the Cambrian, at least for arthropods.     

A venomous arthropod in the Early Cambrian Sea

A number of recently evolved animals possess poison glands for feeding and/or defense.However,examples of such animals are rare in the fossil record.We report a fossil arthropod Isoxys curvirostratus from the Early Cambrian Chengjiang biota of China.This species is regarded as the oldest known venomous arthropod based on the presence of venomous glands in its head region.The adult animal is 2-5 cm long and the body is covered entirely with a carapace.The presence of large stalk eyes and a pair of stout grasping appendages with a terminal spine suggest it was raptorial.Interestingly,the two pear-shaped,three-dimensionally preserved objects that are present in the head region and at the base of the grasping appendages closely resemble the venom glands of some living arthropods in size,shape,and position.These features indicate that the presence of venomous predators could date back 520 million years.Furthermore,our observations suggest that the feeding strategies and organs adapted for this purpose had already reached a high level of diversity and anatomical sophistication in the Early Cambrian ecosystems.     

Possible developmental mechanisms underlying the origin of the crown lineages of arthropods

The extraordinarily preserved, diverse arthropod fauna from the Lower Cambrian Maotianshan shale, central Yunnan (southwest China), represents different evolutionary stages stepping from stem lineages towards crown arthropods (also called euarthropods), which makes this fauna extremely significant for discussion of the origin and early diversification of the arthropods. Anatomical analyses of the Maotianshan shale arthropods strongly indicate that     

A Cambrian micro-lobopodian and the evolution of arthropod locomotion and reproduction

The evolutionary success of arthropods, the most abundant and diverse animal group, is mainly based on their segmented body and jointed appendages, features that had evolved most likely already before the Cambrian. The first arthropod-like animals, the lobopodians from the Early Cambrian, were unsclerotized and worm-like, and they had unjointed tubular legs. Here we describe the first three-dimensionally preserved Cambrian lobopodian. The material presented of Orstenotubulus evamuellerae gen. et sp. nov. is the smallest and youngest of a lobopodian known. O. evamuellerae shows strikingly detailed similarities to Recent tardigrades and/or onychophorans in its cellular-structured cuticle and the telescopic spines. It also shows similarities to other, longer known lobopodians, but which are ten times as large as the new form. These similarities include the finely annulated body and legs, which is characteristic also for Recent onychophorans, and paired humps continuing into spines situated dorsally to the leg insertions, a feature lacking in the extant forms. The morphology of O. evamuellerae not only elucidates our knowledge about lobopodians, but also aids in a clearer picture of the early evolution of arthropods. An example is the single ventral gonopore between a limb pair of O. evamuellerae, which indicates that a single gonopore, as developed in onychophorans, tardigrades, pentastomids, myriapods and insects, might represent the plesiomorphic state for Arthropoda, while the paired state in chelicerates and crustaceans was convergently achieved. Concerning life habits, the lateral orientation of the limbs and their anchoring spines of the new lobopodian imply that early arthropods were crawlers rather than walkers.     

Sophisticated particle-feeding in a large Early Cambrian crustacean

Most Cambrian arthropods employed simple feeding mechanisms requiring only low degrees of appendage differentiation. In contrast, post-Cambrian crustaceans exhibit a wide diversity of feeding specializations and possess a vast ecological repertoire. Crustaceans are evident in the Cambrian fossil record, but have hitherto been known exclusively from small individuals with limited appendage differentiation. Here we describe a sophisticated feeding apparatus from an Early Cambrian arthropod that had a body length of several centimetres. Details of the mouthparts resolve this taxon as a probable crown-group (pan)crustacean, while its feeding style, which allowed it to generate and handle fine food particles, significantly expands the known ecological capabilities of Cambrian arthropods. This Early Cambrian record predates the major expansions of large-bodied, particle-handling crustaceans by at least one hundred million years, emphasizing the importance of ecological context in driving adaptive radiations.     

An armoured Cambrian lobopodian from China with arthropod-like appendages

Cambrian fossil Lagerst?tten preserving soft-bodied organisms have contributed much towards our understanding of metazoan origins. Lobopodians are a particularly interesting group that diversified and flourished in the Cambrian seas. Resembling 'worms with legs', they have long attracted much attention in that they may have given rise to both Onychophora (velvet worms) and Tardigrada (water bears), as well as to arthropods in general. Here we describe Diania cactiformis gen. et sp. nov. as an 'armoured' lobopodian from the Chengjiang fossil Lagerst?tte (Cambrian Stage 3), Yunnan, southwestern China. Although sharing features with other typical lobopodians, it is remarkable for possessing robust and probably sclerotized appendages, with what appear to be articulated elements. In terms of limb morphology it is therefore closer to the arthropod condition, to our knowledge, than any lobopodian recorded until now. Phylogenetic analysis recovers it in a derived position, close to Arthropoda; thus, it seems to belong to a grade of organization close to the point of becoming a true arthropod. Further, D. cactiformis could imply that arthropodization (sclerotization of the limbs) preceded arthrodization (sclerotization of the body). Comparing our fossils with other lobopodian appendage morphologies--see Kerygmachela, Jianshanopodia and Megadictyon--reinforces the hypothesis that the group as a whole is paraphyletic, with different taxa expressing different grades of arthropodization.     

Sonic hedgehog function in chondrichthyan fins and the evolution of appendage patterning

The genetic mechanisms regulating tetrapod limb development are well characterized, but how they were assembled during evolution and their function in basal vertebrates is poorly understood. Initial studies report that chondrichthyans, the most primitive extant vertebrates with paired appendages, differ from ray-finned fish and tetrapods in having Sonic hedgehog (Shh)-independent patterning of the appendage skeleton. Here we demonstrate that chondrichthyans share patterns of appendage Shh expression, Shh appendage-specific regulatory DNA, and Shh function with ray-finned fish and tetrapods. These studies demonstrate that some aspects of Shh function are deeply conserved in vertebrate phylogeny, but also highlight how the evolution of Shh regulation may underlie major morphological changes during appendage evolution.     

鞭角华扁叶蜂预蛹神经内分泌系统的解剖

通过对鞭角华扁叶蜂预肾的解剖,明确屯其神经内分泌系统的构造及其分布。其中枢神经系统由脑、食管下神经节、腹神经索组成,前、中、后脑分化明显。腹神经索包括三对胸神经节、八对腹神经节及纵贯各神经节的神经索,其中各了的分布与叶蜂预肾体节相一致,位于各体节腹板的前缘中部。口道交感神经系统由位于脑前方食道背面的额神经节和位于脑后方的后头神经节组成的神经连锁构成。腹交感神经系统由位于腹神经索各神经节上的横神经组     

Body plan innovation in treehoppers through the evolution of an extra wing-like appendage

Body plans, which characterize the anatomical organization of animal groups of high taxonomic rank, often evolve by the reduction or loss of appendages (limbs in vertebrates and legs and wings in insects, for example). In contrast, the addition of new features is extremely rare and is thought to be heavily constrained, although the nature of the constraints remains elusive. Here we show that the treehopper (Membracidae) 'helmet' is actually an appendage, a wing serial homologue on the first thoracic segment. This innovation in the insect body plan is an unprecedented situation in 250 Myr of insect evolution. We provide evidence suggesting that the helmet arose by escaping the ancestral repression of wing formation imparted by a member of the Hox gene family, which sculpts the number and pattern of appendages along the body axis. Moreover, we propose that the exceptional morphological diversification of the helmet was possible because, in contrast to the wings, it escaped the stringent functional requirements imposed by flight. This example illustrates how complex morphological structures can arise by the expression of ancestral developmental potentials and fuel the morphological diversification of an evolutionary lineage.     

New observations of the lobopod-like worm Facivermis from the Early Cambrian Chengjiang Lagersttte

Facivermis yunnanicus （Hou ＆ Chen, 1989）, from the Early Cambrian Chengjiang Lagerstatte, a worm-like fossil with 5 pairs of tentacles and a perceived shrunken end, has been regarded as related to polychaetes, later it has been variously interpreted as Iobopods, Pentastoma and Iophophorates. Newly discovered complete specimens by the ELI field team show that the taxon has, in addition to the 5 pairs of appendages, a pear-shaped trunk end bearing two or three circles of hooks. Accordingly based on these important morphological characters, reconsideration of its affinities is provided and the taxonomy is remedied herein. Because the five pairs of appendages of Facivermis yunnanicus resemble the appendages of the fore-trunk of Iobopod Miraluolishania （Liu ＆ Shu, 2004）, it seems that they are homologous structures. Therefore, the affinities of Facivermis are regarded here as being close to the Iobopods, in addition, the lobe-like appendages of Facivermis are very crucial to exploring the origin of the appendages of Iobopods and arthropods.     

中国鲎脑神经节的组织学观察

采用组织切片和H-E染色法观察了中国鲎(Tachypleus tridentatus)脑神经节的组织形态结构.研究发现中国鲎的脑神经节分为前脑和后脑.前脑由视神经系统、蕈形体以及蕈形体周围的细胞群组成,后脑由腹面侧后方第一细胞群、侧后方神经纤维网和部分的中央神经纤维网组成.中国鲎的蕈型体比其他节肢动物都发达得多,可能承担着更多的生理功能.研究确认和区分了其脑神经节的细胞群和神经纤维网,同时构建了中国鲎脑神经节的组织形态学图谱,旨在为中国鲎的神经生物学的研究积累资料.     

Skara hunanensis a new species of Skaracarida (Crustacea) from Upper Cambrian (Furongian) of Hunan, south China

The fossils of Orsten-type preservation are as famous as Chengjiang Fauna in the world, but it was not until 2005 that the Orsten-type fossils represented by Skaracarida and Phosphatocopida were first reported to be found in western Hunan, South China. Here, we report the systematic paleontology of all the exquisitely preserved specimens belonging to Skaracarida. They were found at a same horizon of Upper Cambrian (Furongian) in Wangcun section, western Hunan, south China, assigned to a new species Skara hunanensis herein. Skara hunanensis is characterized by small, soft-integumented, marine forms with slender, annulate body; labrum large and ventrocaudally directed; uniramous antennulae; biramous antennae and mandibulae; maxillas and maxillipeds of the same shape; all postantennular limbs join a short cephalic filter apparatus. The body has two tagmata: a cephalon with five pairs of well developed appendages and a trunk composed of 11 ring-shaped conical segments.     

Silurian brachiopods with soft-tissue preservation

'Articulated' rhynchonelliformean brachiopods are abundant shelly fossils, but the direct fossil record of their soft parts was hitherto confined to a single pyritized trace possibly representing a lophophore. Anatomical knowledge of extinct rhynchonelliformeans relies heavily on analogies to extant species; these analogies are untested for stem-group clades. The Silurian Herefordshire (UK) Konservat-Lagerst?tte (about 425 Myr bp) yields exceptionally preserved three-dimensional fossils that provide unrivalled insights into the palaeobiology of a variety of invertebrates. The fossils are preserved as calcitic void in-fills in carbonate concretions within a volcaniclastic horizon, and are reconstructed digitally. Here we describe a stem-group rhynchonelliformean specimen from this deposit; it most probably belongs in the order Orthida. A robust ridged pedicle with distal rootlets is preserved, together with a lophophore and other soft-tissue structures. The pedicle morphology is novel, urging caution in inferring stem-group rhynchonelliformean anatomy from that of crown-group species. Smaller brachiopods are attached to the specimen; these include a probable atrypide, with pedicle and marginal setae preserved.     

稻田与非稻田生境越冬节肢动物群落调查与分析

2008年11月至2009年3月,对清新县双季稻田的冬闲田WF、田埂FR、周围杂草WH以及黑麦草田LL等4种不同生境下的节肢动物越冬群落进行系统调查。结果如下:鉴定出包括昆虫纲8目22科和蛛形纲1目6科共33种节肢动物。其中,植食性昆虫16种,捕食性昆虫4种,寄生性昆虫3种,中性昆虫3种,蜘蛛类7种。闲田和杂草均以蜘蛛类为主要优势种类,而田埂和黑麦草田分别以半翅目和双翅目昆虫为最优势类群。Shannon-Wiener多样性指数(H)和Pielou均匀性指数(J)以田埂的最高,黑麦草田最低;优势度指数(C)以黑麦草田高,田埂最低。杂草地与黑麦草田Jaccard相似性指数(q)最高,而冬闲田与田埂的相似性指数最低。     

Mitochondrial protein phylogeny joins myriapods with chelicerates

The animal phylum Arthropoda is very useful for the study of body plan evolution given its abundance of morphologically diverse species and our profound understanding of Drosophila development. However, there is a lack of consistently resolved phylogenetic relationships between the four extant arthropod subphyla, Hexapoda, Myriapoda, Chelicerata and Crustacea. Recent molecular studies have strongly supported a sister group relationship between Hexapoda and Crustacea, but have not resolved the phylogenetic position of Chelicerata and Myriapoda. Here we sequence the mitochondrial genome of the centipede species Lithobius forficatus and investigate its phylogenetic information content. Molecular phylogenetic analysis of conserved regions from the arthropod mitochondrial proteome yields highly resolved and congruent trees. We also find that a sister group relationship between Myriapoda and Chelicerata is strongly supported. We propose a model to explain the apparently parallel evolution of similar head morphologies in insects and myriapods.